CCRE 2018 Annual Report

Smithsonian Institution

2018 CCRE Annual Report Caribbean Coral Reef Ecosystems • National Museum of Natural History

CCRE Fiscal Year 2018

Table of Contents 4 Carrie Bow Cay Field Station 5 Research Highlights 29 Research Briefs 35 Scientific Publications 37 Visitors and Station Managers 38 Acknowledgements and Photo Credits

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Carrie Bow Cay Field Station

Carrie Bow Cay Field Station has operated on the Belize barrier reef since 1972. The station is open to scientific visitors year-round and offers unparalleled access to coral reef environments, seagrass meadows, and mangrove forests.

This small and highly functional field laboratory boasts a flow-through seawater system, wet and dry laboratory space, full SCUBA facilities, research vessels, and living quarters.

For more information, visit: www.ccre.si.edu

Thalassia Experimental Network: a multi-institutional effort to evaluate herbivory and nutrient enrichment effects on turtlegrass ecosystem function

Justin Campbell, Valerie Paul, Andrew Altieri, James Douglass, Ken Heck, Olivia Rhoades

Seagrasses are foundational species that provide a range of services to ecosystems, including habitat provisioning, food, sediment stabilization, and improved water quality. Both top-down (herbivory) and bottom-up (nutrients) factors impact the growth and production of seagrasses across temperate and tropical regions, yet the relative importance of these processes in controlling seagrass ecosystem function remains controversial, particularly across latitudinal gradients in light and temperature. In 2018, we established the Thalassia Experimental Network (TEN), a seagrass network involving eleven institutions working across thirteen sites. These sites span from Bermuda to Panama and encompass nearly thirty degrees of latitude. One of our sites is at Carrie Bow Cay. In the late spring, we deployed a caging experiment in a shallow seagrass bed at each site. The bed was divided into 50 small plots (0.25 x 0.25m) and each plot received a unique combination of treatments in a factorial design: caging (to exclude natural herbivores), artificial grazing (seagrass cut short with scissors to simulate increased herbivory), and nutrients (added nitrogen and phosphorus). Each experiment will last for one year, after which we will measure a range of seagrass characteristics including biomass, productivity, associated epifauna, sediment composition, and nutrient content. We have already collected some preliminary samples (in late summer), and we are currently in the process of analyzing these samples to detect initial trends. Anecdotally, and in accordance with findings from a preliminary study in 2014, we are witnessing a strong effect of nutrient enrichment, which influences seagrasses via distinct mechanisms across latitudes. Nutrient enrichment reduces seagrass production by increasing epiphyte growth at our northern, sub-tropical sites; while it enhances rates of herbivory on nutrient-enriched seagrass at the tropical sites.

Final results from our network will elucidate the relative importance of top-down versus bottom-up factors in driving the ecological functioning of seagrasses across gradients in light and temperature. Our results will also allow us to better predict the effects of climate change, including rising sea surface temperatures and the tropicalization of seagrass ecosystems across sub-tropical locations.

Research diver at seagrass site near Carrie Bow Cay.

Caribbean Coral Reef Ecosystems | 6

Understanding fish and coral settlement cues

Skylar Carlson, Jennifer Joseph, Zara-Louise Cowan, Jennifer Sneed, Audrey Looby, Lane Johnston, Kieran Cox, Abigail Engleman, Valerie Paul, Danielle Dixson

This project seeks to identify the reef-based chemical cues used by larval fishes and corals to locate appropriate reef habitat. The majority of coral reef fishes are hatched from eggs and enter a planktonic stage. In this stage, they are released into the water column and journey far from the reef. However, at the conclusion of the larval stage, fish must locate suitable habitat from the open ocean to transition into benthic juveniles. Chemical cues carried from the reef into offshore waters through tides and currents are an important mechanism employed by fish to find their way back to the reef. This project seeks to identify the specificity of different cues responsible for guiding these fish home and gain insights into the chemical nature of these cues. In Carrie Bow Cay’s wet lab, we offer larval fish a choice between seawater and seawater containing different chemical cues. The cues are prepared by soaking individual species removed from the reef in seawater for a set period of time (usually one hour).

We have examined algae, adult coral, cyanobacteria, seagrass, sponges, and other fish species as sources for potential attractive and deterrent cues. Preference or avoidance behaviors by fish are tested using a two-choice flume where water flows side by side and the location of the individual larval fish or new recruit is recorded.

In addition to the choice flume, preferences towards specific coral reef components (algae, coral, rubble) were also tested using baby pools set up at the station. Here fish were offered the choice of small patch reefs within the pools, and the time spent associated with each reef was recorded to identify a preference. Coral larvae are tested in much the same way but some species of coral only spawn once a year and actively choose settlement locations for only 48 hours. In the laboratory in Fort Pierce, we extract the same members of the reef that are found to attract or deter larvae in order to find the compounds that might be involved as cues. These extracts are then tested in the field to determine if the molecules that were attractive (or deterrent) have been successfully extracted. These extracts are then fractionated and retested to try to determine if individual molecules can be identified that affect fish behavior. These efforts are the initial steps toward determining what an attractive reef “smells” like to guide knowledge-based conservation and restoration efforts.

This year, we were joined in the field by two Smithsonian Link Fellows who worked on subaims of this project.

Abigail Engleman is working to understand how structure and chemical cues work together to recruit and build healthy reefs. She incorporated coral larval settlement cues on settlement tiles in the laboratory to test the effects of structural complexity and chemistry of coralline algae that are known to facilitate settlement and recruitment of coral larvae (see pages 9-10). Kieran Cox builds 3D models from photographs to measure the complexity of a reef habitat and determine the impact of structural complexity on fish biodiversity. His work will help us better understand reef composition and the characteristics of a diverse and healthy reef.

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Studies on reef fish provide useful insights into the role of chemical cues in settlement.

Assessing synergy of chemical and physical stimuli on coral settlement Abigail Engleman

Coral, an animal that is immobile after its larval stage, must choose a habitat that will give it the greatest chance of long-term survival. Coral make this decision using sensory anatomy to sense abiotic cues, triggering when and where they should settle on the seafloor. Understanding physical and chemical cues that drive this critical life-history stage is fundamental to advancing our understanding of coral reef ecosystems.

Structures, for example, play an important role in facilitating coral larval movement towards the seafloor, and enhance coral’s ability to remain attached once they settle. Research suggests that coral abundance is higher on areas of natural reefs which have small crevices rather than flat surfaces. Perhaps that is because smallscale three-dimensional structures alter water flow, guiding larvae towards the seafloor, while small topographic features provide additional ‘attachment points’ for coral to securely affix themselves to the substrate.

These structural characteristics (among others) help larvae determine whether a habitat is a suitable location to begin their sessile life.

The current research challenges this notion by assessing how different structure-types affect coral recruitment, and whether structures synergistically enhance coral survival, when combined. With funding from the National Geographic Society, this research aims to advance fundamental knowledge about coral life-history, and uncover more information about how coral choose a permanent home. Field experiments for this study began July 2018, when we deployed settlement tiles across three reef habitats off Carrie Bow Cay, Belize. These tiles were specially designed with features to test whether multi-scale structural complexity influences coral recruitment. After successful deployment, tiles must remain on the reef for months, allowing time for coral to settle and establish residence on the structures. Thus, we will remove the tiles in 2019 to analyze coral recruitment back at the Carrie Bow Cay laboratory.

Increasing evidence suggests coral reefs’ structural complexity has decreased in recent years. Though still ongoing, this project will likely yield important information regarding coral recruitment preferences, which influence long-term sustainability of reef structure. Determining what characteristics coral seek in their foreverhome will advance our understanding of coral reef ecosystems and their fate. These findings are essential for developing effective management, conservation, and restoration strategies for coral reefs around the world.

Coral settlement tile with model coral fragment.

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Monitoring the effects of global change on marine biodiversity and ecosystem function at Carrie Bow Cay, Belize Maggie Johnson This study is characterizing present-day ecosystem function on coral reefs of Carrie Bow Cay, Belize, specifically early successional community structure and net carbonate accretion. These are important ecosystem properties that indicate the potential for coral reefs to continue growing by accreting calcium carbonate. This research contributes to the Smithsonian’s Marine Global Earth Observatory (MarineGEO) monitoring program, and the results from Carrie Bow Cay will be directly compared to paired experiments in Bocas del Toro and Coiba, Panamå. This approach will provide important information that can inform strategies to manage and monitor the effects of global change on coral reef ecosystems.

The primary objective of this research is to monitor early successional community structure, biodiversity and net ecosystem accretion using calcification accretion units (CAUs). These data will provide information on the current status of coral reefs of Carrie Bow Cay, Belize, how ecosystem properties are changing over time and how they compare to other coral reefs in the Caribbean and Pacific. CAUs were initially deployed at 6 sites around Carrie Bow Cay in 2016. These units were collected in 2017, and a new set were redeployed. The 20172018 units were collected during the 2018 research trip to Carrie Bow Cay, marking the second consecutive year of monitoring and a new set (2018-2019) were deployed. In September 2018, CAUs were retrieved from 6 sites (5 at each site), for a total of 30 units. Each of the units was disassembled at the Smithsonian research station on Carrie Bow Cay, photographed, and dried. Dried tiles were returned to the Smithsonian Marine Station in Fort Pierce, Florida and are currently being analyzed for the amount of calcium carbonate deposited on each tile. Photographs of tiles are also being analyzed to determine the percent cover of dominant functional groups. Data are still being collected from the units that were deployed from 2016-2017, and these tiles (2017-2018) will take many more months to be fully processed.

Similar units are also being used to assess calcium carbonate accretion on coral reefs in Panama. Thus the data from Carrie Bow Cay contributes to a larger data set that provides insight into the environmental factors that contribute to reef growth (calcium carbonate accretion) on case study reefs in the Caribbean. In addition to providing baseline data on reef accretion, they provide a metric that can be monitored over time to see how reefs respond to changes in the environment.

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The author deploying and retrieving calcium accretion units (CAUs) near Carrie Bow Cay.

Testing the sponge-loop hypothesis for Caribbean coral reefs

Joseph R. Pawlik, Steven E. McMurray and Christopher M. Finelli

Sponges dominate Caribbean reefs now that reef-building corals have been declining for decades. Sponges feed by filtering huge volumes of seawater, providing a mechanism for transferring organic carbon back to the benthos as they turn-over the water column. A new theory has been proposed about benthic-pelagic coupling on coral reefs called the “sponge-loop hypothesis” that is potentially the most important new concept in marine ecology in many years, because it seeks to explain Darwin’s Paradox: how do highly productive and diverse coral reefs grow in desert-like tropical seas? Similar to the famous microbial-loop of the 1980s, the sponge loop hypothesis proposes that sponges on coral reefs absorb the large quantities of dissolved organic carbon (DOC) exuded by carbon-fixing seaweeds and corals and return it to the benthos as particulate organic carbon (POC) in the form of shed cellular detritus. But does the sponge-loop hypothesis apply to the sponge species that make up the greatest biomass and process the most seawater on coral reefs? On the fore-reefs of Carrie Bow Cay, we investigated the components of the sponge-loop hypothesis for common barrel, vase and tube-forming sponge species that span a range of associations with microbial symbionts, from high microbial abundance (HMA) to low microbial abundance (LMA) in the sponge tissue. We used InEx techniques, velocimetry, and flow cytometry to determine whether each species consumes DOC and produces cellular detritus (POC). Then, for species that consume DOC, we used the same techniques in manipulative experiments that augment the amount of DOC to determine the types of DOC consumed by sponges. The results of this project will be important in determining how DOC is transferred from the water-column to the benthos as part of the carbon cycle in coral reef environments where the effects of global climate change and ocean acidification may be tipping the competitive balance toward non-calcifying organisms, such as sponges.

Researchers from the Pawlik lab are examining the role sponges play in recycling organic carbon on coral reefs.

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What factors control distribution and abundance of sponges living in coral reef, seagrass meadow, and mangrove habitats?

Janie Wulff

Although experiments have demonstrated many key ecosystem roles for sponges on coral reefs and in mangroves, relatively little is known about population and community dynamics of sponges. Learning what controls sponges is important because the roles they play are not duplicated by other groups of organisms, and many of these roles influence health of corals, mangroves and their ecosystems in dramatic ways. Variation among sponge species in roles played and vulnerabilities to environmental challenges means that identification must be to species and that abundance must be evaluated by measuring volume. Thus their very high species diversity and unusual shapes have impeded inclusion of sponges in long-term monitoring programs. Our lab has been maintaining sponge census plots in coral reef, seagrass meadow, and mangrove sites near Carrie Bow Caye for many years, and results of repeated censusing have often been surprising. Because sponges vanish when they die, mortality events can only be recorded when in progress or if they occur at a site where sponges have been previously censused, resulting in chronic underestimates of sponge mortality. Mass mortality of coral reef sponges at the Blue Ground Range, due to a dense phytoplankton bloom in 2011, was revealed by our censusing. A primary focus of field work at Carrie Bow Caye in 2018 was continuation of the coral reef censusing to document both recovery from that 2011 loss of more than 2/3 of the sponge biomass, as well as documenting subsequent mortality events. Continued censusing of sponges on mangrove prop roots at Twin Cayes allowed detailed documentation of a mass mortality event in April/May 2018 that was caused by thick Sargassum mats and resulted in loss of the protective function of sponges against isopods that diminish mangrove survival by burrowing into prop roots as well as a nutrient exchange mutualism between sponges and mangroves.

Photos of complete mortality on mangrove prop root sponge Tedania ignis (left) and almost complete mortality in Dysidea etheria (right). The blue tissue is what remains of the living sponge. Both species were affected by large Sargassum mats retained among mangrove roots at Twin Cays.

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Sponges play important roles in a variety of marine ecosystems.

Is hybridization among threatened Caribbean coral species the key to their survival or the harbinger of their extinction?

Nicole Fogarty, Megan Bock, Kelly Pitts, Morgan Hightshoe While coral cover has declined worldwide, Caribbean acroporids have experienced some of the highest mortality. Population losses of the corals Acropora cervicornis and A. palmata have reached up to 90% since the 1970s and have been primarily attributed to disease and the loss of symbiotic algae (i.e., coral bleaching) from thermal stress. As global sea surface temperatures (SST) rise, mass bleaching events are increasing in frequency and severity, posing a serious threat to coral reefs. To predict future population success of the Caribbean acroporids, including their hybrid, A. prolifera, we conducted a series of thermal tolerance experiments at Carrie Bow Cay, Belize.

Fragments of A. cervicornis, A. palmata, and A. prolifera collected from local reefs were relocated to an underwater nursery established by the Fogarty Laboratory at Nova Southeastern University. These fragments were grown in the nursery for over a year prior to experimentation to ensure they were exposed to identical environmental conditions and possessed the same symbiotic algae. In May 2018, fragments were placed into acrylic aquaria on the island, where they were subject to either increased or ambient water temperatures for a 72-hour period. During this time, each coral’s photosynthetic efficiency was measured daily. Oxygen consumption and production of corals was also measured using custom-made respiration chambers. Following the 72-hour exposure, samples were taken to measure chlorophyll a, zooxanthellae (algae), and protein concentrations in the coral tissues.

Regardless of species, corals exposed to high water temperatures had significantly lower concentrations of zooxanthellae, proteins, and chlorophyll a when compared to those kept in ambient conditions. Temperature stressed corals also were unable to photosynthesize during respiration trials. Based on bleaching prevalence, A. cervicornis visually appeared to be more tolerant of high water temperatures than A. palmata and the hybrid, and also had significantly higher total and host protein concentrations following exposure. Once analyzed, these results will be used to better understand differences in the Caribbean acroporids and how populations may change with future ocean conditions.

Nova Southeastern University graduate students, Morgan Hightshoe (left) and Megan Bock conducting experiments with Acropora fragments at Carrie Bow Cay.

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Citizen Science GIS: Open Reef Timothy L. Hawthorne and Nick Altizer Open Reef is a research initiative through Citizen Science GIS, a 2017 Esri Special Achievement in GIS Award winner, at University of Central Florida that changes the way science and society see and explore vulnerable island environments around the world through drone mapping, open data, storytelling, and citizen science.

Through the use of low-cost, easily accessible drone technology we create high-resolution, open-source imagery of the Belize islands. Features such as island boundaries, structures, docks, and seawalls are then analyzed to understand developmental changes and vulnerabilities on each island. All the while we are interacting with and involving local communities to learn more about the changes that are occurring and to share their stories through the process of spatial storytelling. This summer, we interviewed 42 local residents, fisherman, guides, and resort workers to learn about their experiences with climate adaptation, island changes, and resiliency.

Open Reef worked again this summer in Belize, including a weeklong mapping project at Carrie Bow. This year’s work at Carrie Bow was primarily funded by a U.S. National Science Foundation Research Experiences for Undergraduates Site grant and additional support from MarineGeo. Open Reef mapped important reef and island features on the trip, ranging in size from 13 acres to 1100 acres. With our drone mapping capabilities we are able to collect imagery at a resolution of a few centimeters, providing a much higher resolution and level of detail than most freely available satellite imagery in Belize. To date, our work has been used by the field station, Southern Environment Association, Coastal Zone Management Institute and Authority, the Village of Hopkins, Fragments of Hope, and other organizations in Belize.

Open Reef also fosters positive education outcomes by connecting educators from different disciplines to support Belize and U.S. student development. During our summer work in Belize, the team held our annual Hopkins Village Citizen Science GIS Youth Academy as Miss Bertie’s Community Village Library. Over 150 youth from the village engaged in drone mapping, mini-drone flights, and online mapping and fieldwork. The program ended with a boat trip and tour supported by Hamanasi Resort to Carrie Bow Caye, drone flights in the South Water Marine Reserve, and a snorkel trip. Open Reef aims to support future scientists in Belize and beyond. Our open data platform allows for shared partnership opportunities, improved data quality, and community engagement and input. To find out more please visit www.citizensciencegis.org/openreef or contact Dr. Timothy Hawthorne at timothy.hawthorne@ucf.edu.

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High resolution drone image of Carrie Bow Cay Field Station.

Paragoniolithon solubile

Titanoderma bermudense

Paragoniolithon acretum

The Diversity of Encrusting Red Algae Raphael Ritson-Williams

Throughout the Caribbean the diversity and distribution of crustose coralline algae (CCA) are poorly known.

These encrusting red algae play multiple ecological roles on coral reefs, yet their taxonomy remains poorly understood due to few morphological features that are difficult to observe without high powered microscopes.

This research was designed to catalog the diversity of CCA found at Carrie Bow Cay along the barrier reef of Belize.

While at Carrie Bow Cay, 35 individual specimens of CCA were photographed live, collected, dried and preserved for morphologic and genetic analysis. Samples were collected from a variety of habitats ranging in depths from 0 to 60 feet. This collection included 13 species of the 27 species that are thought to occur in the Caribbean. The dried samples were brought back to the California Academy of Sciences in San Francisco, where every specimen was morphologically characterized under the scanning electron microscope to identify the species. Using small fragments of the same dried samples, they were extracted for their DNA and sequenced using multiple genetic markers. In this way I have built a database of morphology and DNA for every specimen.

I will continue to work on the genetics of these specimens and will make the phylogeny open access as soon as I have completed the analysis. This genetic database will serve as an important reference so that other researchers working on CCA can determine which species of CCA they have by sequencing a small piece of their sample.

Additionally, I have been building a series of species pages to describe live characters of each species so that researchers can begin to identify CCA species in the field. I have completed 5 of these field guides and they are available open access on my website: www.raphswall.com/cca. As I continue to build the morphological and genetic data for this project more species pages will be made available on my website.

Representatives of CCA species found near Carrie Bow Cay.

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CARICOMP: Long-term loss of reef structural complexity on the forereef at Carrie Bow Cay Karen Koltes and John Tschirky

The Caribbean Coastal Marine Productivity (CARICOMP) program was initiated in 1990 as a basinwide cooperative network of marine laboratories.

The goals were to monitor the physical environment using standardized methods and document trends in the structure and functioning of coral reefs, seagrasses and mangroves.

CARICOMP Monitoring of coral reefs using the CARICOMP protocols began at Carrie Bow Cay in late 1993. Per CARICOMP methodology, a study area was chosen on the inner forereef slope east of Carrie Bow Cay that was “representative” of the main coral reef community in the depth range of 10-13m. This zone is dominated by columnar coloFig. 1. Coral bleaching, primarily of O. annularis colonies, at nies of the star coral, Orbicella annularis, one of the the CARICOMP site, December 1995. major reef-building corals in the Caribbean. When the inner forereef slope was first surveyed in the 1970’s by Smithsonian scientists, however, it was dominated by another of the major reef building corals, the staghorn coral, Acropora cervicornis. By 1984, most of the A. cervicornis had died off from white band disease (Acroporid white syndrome) and has not recovered in the almost 40 years since.

Following the CARICOMP protocol, the study area on the inner forereef slope was separated into two sub-areas. The northern sub-area was selected as it overlaps the original 1970’s Smithsonian surveys. Depths range from about 12-15 m. The southern sub-area was chosen for being a large area with a nearly uniform depth of about 13-14m. At each sub-area, five permanent, 10m transects were established using stainless steel poles.

Transects are surveyed using the linear chain method in which a light chain is draped over the substrate underneath a measuring tape stretched between the two transect poles. The number of links of each organism or substrate type is recorded and organisms are identified to species where possible. In addition to providing data on benthic community structure (e.g., percent coral cover), the chain method also provides an estimate of rugosity, a measure of the three-dimensional structural complexity of a reef. Rugosity is an important ecological parameter as areas of highly complex architecture provide more habitat for reef fish, corals, algae and motile and sessile invertebrates. Reef flattening (loss of rugosity) is among the major problems currently faced 23 | Caribbean Coral Reef Ecosystems

by Caribbean coral reefs. Rugosity is calculated as the ratio of the total chain length (total number of links) draped over the substrate divided by the horizontal distance (10m).

The 10 CARICOMP reef transects have been surveyed annually or semi-annually since 1993 except for a 2-year period following the destruction of the field station by a fire in December 1997. Significant bleaching of corals, especially of O. annularis, at Carrie Bow Cay was first observed in December 1995 (Fig. 1). This was followed in 1997 and 1998 by the El Nino Southern Oscillation that triggered severe bleaching of coral reefs around the world, including at Carrie Bow Cay (Fig. 2). In late October 1998, Hurricane Mitch, a Category 5 hurricane, struck Central America. Mitch was one of the strongest and most damaging storms ever to hit

Fig. 2. Bleached colonies of O. annularis on the inner forereef slope south of the CARICOMP site.

Bleaching began in the summer of 1998, triggered by record high temperatures associated with the 1997-1998 El Nino. The photo was taken about 10 days after passage of Hurricane Mitch in November 1998.

CARICOMP, continued

the Caribbean. At its height on October 26 and 27, the hurricane had sustained winds of 180 mph and dumped more than a meter of rain over Central America, resulting in the deaths of 10,000 – 20,000 people. Mitch was the first hurricane to hit Belize after a nearly two-decade period of little-to-no hurricane activity.

A key observation from the long-term monitoring program is that the inner forereef slope in the 10-15m depth range suffered significant damage following Hurricane Mitch. Rugosity declined by about 25%, from an average of 1174 links/transect between 1993 and 1997 (the last survey before the fire) to a low of 887 links/ transect by 2003 (Fig. 3). Moreover, there has been little recovery of reef complexity in the past two decades.

Fig. 3 The CARICOMP transect about 10 days after passage of Hurricane Mitch. A broken column of O. annularis can be seen on the substrate under the yellow slate. Note that the substrate is composed almost entirely of the skeletons of A. cervicornis. Once dominant in this zone, it died off in the 1980’s from white band disease and has not recovered.

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The immediate loss of rugosity following Hurricane Mitch was the result primarily of the scouring of the substrate and breakage of the large (~1m high by ~1 m diameter) colonies of O. annularis.

The lack of recovery of rugosity in the intervening two decades can be attributed to a number of factors, including: 1) recruitment of faster growing reef components that contribute to rugosity, such as sponges and the lettuce coral, Agaricia tenuifolia, that has been offset by the continued loss of colonies of O. annularis (Fig. 4); 2) the abundance of skeletons of A. cervicornis following the 1980’s die-off (Fig. 2) created an unstable substrate which inhibited recruitment; 3) an increase in the frequency of storms and bleaching (and subsequent disease outbreaks) in the 2 decades following Mitch compared to the two decades preceding it; 4) bioerosion in excess of growth of O. annularis and other reef-building corals; and 5) the slow growth rate (~ 6-10 mm/yr) of O. annularis will require decades for new colonies of this species to achieve the size of those lost, assuming no other stressors.

The 25-year record of coral reef surveys showing loss of rugosity on the inner forereef slope is consistent with other findings in

the region of degradation and flattening of reef structure. Loss of reef structural complexity is likely to have serious consequences for reef biodiversity, ecosystem functioning and the environmental services that reefs provide. Results of the CARICOMP program demonstrate the value of long-term monitoring in being able to document the ways in which major environmental perturbations shape current reef conditions even after two decades on the barrier reef around Carrie Bow Cay.

Fig. 4. Mean rugosity measurements of the 10 permanent CARICOMP transects at Carrie Bow Cay. Rugosity dropped sharply following Hurricane Mitch and has failed to recover in the past 20 years.

Impacts of microbes and plastics in river discharge on coral health and algal phase shift mechanisms on coral reefs in Belize Matthew Hoch, Emily Smith, Mark McNab, Katelin Catching, Ashlyn Borel, Hayden Henslee, Craig Nelson, Linda Wegley

Coral reefs support tremendous biodiversity and essential ecological services, yet multiple human impacts over recent decades are resulting in declining coral health, increased benthic fleshy macroalgal dominance, and biodiversity loss. The mechanisms of coral decline and macroalgal replacement are hypothesized to involve positive feedback interactions among macroalgae, dissolved organic matter (DOM), pathogenic microbes and coral disease. Plastics pollution from rivers also increases the incidence of coral disease. This project experimentally tested responses of microbial communities (microbiomes) in plankton and corals exposed to conditions of pulsed river discharge (terrestrial DOM, nutrients, sediment, terrestrial microbes and plastic pollution from the watershed) in the presence and absence of benthic macroalgal DOM exudates. We tested the synergistic effects of river water and macroalgal DOM exudates on plankton and coral microbiomes, the fate biofilm microbiome composition on river-borne plastic debris it transports from river to reef, and the mechanisms of plastic debris from rivers in causing coral disease.

Dissolved organic matter, disease, algae, microorganism (DDAM) positive feedback loop model of coral loss incorporating the proposed exacerbating impacts of large river discharge events.

Belize watersheds adjacent to the South Water Caye Marine Reserve are characterized by steep topography of headwaters in the Maya Mountains and predominance of agriculture and human settlement in mid and lower reaches. Heavy sustained rain events in these watersheds can result in “pulsed� discharge of nutrients, sediments, terrestrial DOM and microbes into coastal waters, which can reach the barrier reef. How these river constituents impact microbiomes of plankton and corals at the reef was explored under conditions with and without excess macroalgal DOM exudate production. Seawater cultures were amended with 10% river water (unfiltered or 0.22 um filtered) and algal exudate (from Dictyota sp. or Galaxaura sp.) in 2x2 factorial design experiments.

The results of these experiments are still awaiting microbiome analysis.

Terrestrial microbes, including pathogens, on plastics in river discharge may be transported in plume water to corals of the Mesoamerican Barrier Reef, Belize, where they may be involved in coral disease initiation. The 27 | Caribbean Coral Reef Ecosystems

proposed approach to test the stability of the plastic biofilm microbiome composition during its transport from river to reef was to sample plastics in the North Stann Creek plume of a major discharge event during the rainy season. However, in the absence rain in late June 2018 an experiment was performed to simulate the exposure of river plastic microbiomes to changes in water quality and bacterioplankton microbiomes as river water mixes with seawater. Over a period of four days, water quality was monitored and plastic pieces collected at several time intervals. DNA was extracted from the plastic samples and bacterioplankton of river water and seawater and a targeted metagenomic analysis of 16SrDNA was performed. Taxonomy and beta-diversity analysis reveal a progressive change, whereby some river microbes were lost and some marine microbes colonize the plastics.

However, many river microbes were able to survive the transition to marine conditions at the Mesoamerican Barrier Reef, including taxa of known coral diseases. Therefore, plastic pollution discharged from watersheds serves as an abiotic-vector for river microbe transport, including some pathogenic bacteria, where they may play a role in impacting coral health on the Mesoamerican Barrier Reef.

Although the incidence of coral disease has been demonstrated to be greater for coral colonies in contact with marine plastic debris, the potential mechanisms for plastics initiation of coral disease is under-studied. Plastics in contact with coral could directly transfer their biofilm microbes, including pathogens, to the coral microbiome. Alternatively, plastic contact could facilitate abiotic conditions favoring growth of opportunistic pathogens already present in seawater or coral mucus.

To test these two hypotheses, colonies of Porites astreoides were incubated in contact with either microbial colonized plastic from a river (direct transmission test) or clean plastic (indirect facilitation test), and the microbiomes of the seawater, coral mucus, and plastics biofilms compared to those of coral and seawater controls. Microbiomes of mucus and seawater were similar in coral controls and coral with clean plastic treatments, and the microbiome that colonized the clean plastic was most similar to that of the seawater. Some change in the river colonized plastic microbiomes occurred, likely due to exposure to seawater salinity. However, the mucus microbiomes of corals in contact with colonized plastic changed significantly and coral colonies became symptomatic of white plague disease (WPD). Specifically, the relative abundance of Rhodobacteraceae and Flavobacteraceae of mucus microbiomes increased 3-fold compared to control Experiment to test the mechanism and clean plastic treated corals, which is indicative the WPD in other coral spe- of coral disease initiation by plastic cies. The negative effects of land-derived plastic debris on coral mucus microbi- debris. Note, strips of clean or river omes and colony health supports that plastics can directly transmit coral disease. microbe colonized black HDPE film covers a portion of the Porites astreoides colonies.

Carbon and Sulphur metabolism in stilbonematid symbioses Sylvia Bulgheresi and Gabriela Paredes

Oxygen is absolutely required to sustain eukaryotic life. Nonetheless, in marine sediments, such as those surrounding the island of Carrie Bow Cay, this element becomes scarce within the first few centimeters. It is in this extreme environmentv that the symbiotic nematode Laxus oneistus thrives. During our 2018 field trip, we performed experiments to assess the effect of oxygen on the metabolism of L. oneistus and on that of the bacterial symbiont which ensheaths its body. More specifically, we incubated batches of symbiotic nematodes in glass vials containing either oxic (190-200 µm O2), hypoxic (<60 µm O2) or anoxic (0 µm O2) sea water for up to 72 h. Anoxic incubations were achieved with the aid of a polyethylene glove bag filled with nitrogen and the concentration of dissolved oxygen inside the incubation vials was monitored throughout the experiments. Upon incubation, samples were preserved for transcriptomics, proteomics and lipidomics to be performed in Austria, Germany and the UK. We believe that these analyses will reveal the survival strategies L. oneistus evolved to withstand anoxia and whether its microscopic friend is giving it a helping hand.

The nematode worm, Laxus oneistus, can be found inhabiting marine sediments near Carrie Bow Cay. Bacterial symbionts ensheath the body of the worm

29 |Caribbean Coral Reef Ecosystems

Mangrove root ascidian ecology Alexander Strawhand and Janie Wulff

Mangrove root communities in Belize are complex habitats classified by a colorful and rich diversity of marine invertebrates that can differ wildly even between roots directly adjacent to one another. At Twin Cays, the most conspicuous member of the communities are sponges in many areas but the pervasiveness of compound Didemnid ascidians suggests they may significantly contribute to the underlying community dynamics. One of the common ascidians, Didemnum conchyliatum, has a tough network of internal calcareous spicules and often occurs in small orange patches or at the tips of roots. Another common ascidian, Diplosoma glandulosum, has no spicules but seems to occupy large areas on roots, with single colonies monopolizing entire roots at times. Our research at Carrie Bow Cay largely focused on observing growth and survival for individual ascidian colonies in order to help understand their roles in community dynamics.

The ascidian Didemnum conchyliatum is commonly found growing on mangrove roots at Twin Cays, Belize.

Sponge-mesofauna symbioses Kate Hill and Janie Wulff

Sponges form symbioses with a wide array of mesofauna including polychaetes, crustaceans, brittle stars, and bivalves. The mesofauna associated with sponges can be incredibly abundant and diverse. Ribeiro et al. (2003) found 2,235 individuals representing 75 invertebrate species from 19 specimens of the sponge Mycale microsigmatosa. My goals with this research were twofold 1) to document the spatial variation of mesofaunal communities associated with Tedania ignis and Tedania klausi and 2) to examine the relationship between Mycale microsigmatosa and serpullid worms. I compared the mesofaunal communities associated with the sponges Tedania ignis and Tedania klausi in seagrass beds and mangroves near Carrie Bow Caye with those in the Florida Keys and Panama, in order to examine the relative influence of host sponge, habitat, and geographic location on mesofaunal communities. The data suggest that many Mycale microsigmatosa with symbiotic mesofauna have species tube-dwelling polychaete worms. specific host associations and that mesofaunal communities are distinct at both the habitat and geographic scale. With respect to the relationship between M. microsigmatosa and serpullid worms, I began monitoring M. microsigmatosa individuals with and without serpullid worms and will follow the marked individuals through time.

Tedania klausi in seagrass meadow.

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Assessing cryptic diversity of Caribbean Ircinia Joseph Kelly

Microbiomes are quickly gaining recognition as ubiquitous and integral components of multicellular organisms. In the sponge genus Ircinia, microbes govern the host’s interface with the environment by performing nutrient cycling, production of toxic defensive compounds, and photosynthesis, activities that have cascading effects throughout the ecosystem. Evolutionary responses to this symbiosis also abound as Ircinia and its microbiome, collectively termed a holobiont, exhibit several hallmarks of metabolic integration including gene losses in the microbes’ genomes, transfer of nutrients from the microbes to the host, and transmission of microbes between host generations through the sponges’ larvae. Thus, microbes in Ircinia hold the potential to influence the quality of the surrounding habitat and the evolutionary histories of their hosts.

This study uses host and microbial DNA sequence data to bring ecological relevance to species delimitation in the sponge genus Ircinia, a group whose taxonomy has proven difficult to study on the basis of host morphology alone. During the author’s visit to the Carrie Bow Cay Field Station in the summer of 2018, specimens belonging to at least three putatively new Ircinia species were collected and are currently undergoing preparation for DNA sequencing.

In organisms where evolutionary impacts of microbiomes are likely prominent, efforts to distinguish among independently evolving lineages should consider both the hosts and their microbial consortia. This approach acknowledges the ecological context of species delineations and provides a means for identifying species boundaries in microbe-rich clades that otherwise lack taxonomically informative morphological characters, such as many groups of corals and sponges.

Given the role sponge microbiomes play in ecological functioning, understanding sponge biodiversity is essential to reliably assess coral reef health and resilience. My study is contributing to this effort by generating clear descriptions of new sponge species that have readily interpretable ecological relevance. These species descriptions will allow biodiversity estimates to be adjusted to reflect true species counts and better inform natural resource management decisions involving Caribbean coral reefs.

A sponge from the genus Iricinia growing on mangrove roots near Carrie Bow Cay, Belize.

2018 TMON field campaign

Scott Jones, Valerie Paul, Maggie Johnson, Scott Ling, David Branson, Michael Goodison, Kieran Cox, Bethany Gaffey, and Maggy Benson

Now in its fourth year, the annual MarineGEO Field Campaign collects critical biodiversity data that contribute to a global network of marine research sites. Belize serves as a critical node in this network due to the high biodiversity found in the region as well as the wealth of historical data collected there. The aim of MarineGEO is to create a long-term, open-source ecological database of coastal marine habitats and biodiversity to determine habitat trajectories over time and inform research direction and management decisions. To do this, we 1) monitor common marine habitat condition and extent, 2) quantify biodiversity in each habitat, and 3) use standardized experimental approaches to quantify ecosystem processes that drive observed patterns. Our specific primary Researchers conducted intensive underwater surveys near Carrie questions are: 1) What is the condition of Bow Cay. major marine habitats around Carrie Bow Cay? 2) How much biodiversity is associated with each habitat type? 3) How do ecological processes (e.g.: consumption rates) vary among habitats?

In the span of a very busy two weeks, researchers conducted a rigorous set of protocols to examine each important tropical habitat surrounding Carrie Bow Cay, including mangroves, seagrasses, patch reefs, sand flats, and fore reef. Intensive assessments of mangroves and seagrasses measure productivity, growth, and biomass, while an exhaustive survey of fish and invertebrate diversity is conducted at each habitat using Reef Life Survey protocols (www.reeflifesurvey.com). Important ecological processes like herbivory and predation are measured with fish feeding assays deployed at each habitat. The data generated from these efforts are impressive: thousands of feeding assays have been recorded and hundreds of fish and invertebrate species have been documented. These projects play an important role in advancing MarineGEOâ&#x20AC;&#x2122;s efforts to monitor coastal ecosystems. Additionally, the scientific team worked to engage with the public during the campaign by hosting a Facebook Live event with the Smithsonianâ&#x20AC;&#x2122;s Ocean Portal (www.ocean.si.edu) and by hosting a live presentation for community members and University of Belize students in Dangriga, Belize.

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CCRE South Water Caye Marine Reserve reef assessment

Scott Jones, Zach Foltz, Randi Rotjan, Leah Harper, and Alicia Volllmer

This year marks the eigth year Caribbean Coral Reef Ecosystems program (CCRE) staff and collaborators have conducted the South Water Caye Marine Reserve (SWCMR) Reef Assessment Program. At over 100,000 acres, SWCMR is one of the largest marine reserves in Belize and encompasses the waters around the Smithsonian’s Carrie Bow Cay Field Station. The reef assessment project aims to identify effects of the no-take conservation zone around Carrie Bow Cay on the recovery of fish and coral populations. In June 2011, permanent transects were established inside (12 transects) and outside (12 transects) the area’s boundary and have been surveyed biannually since then. This amounts to over 336 unique surveys on the reefs around Carrie Bow Cay. Reef monitoring efforts typically measure the diversity, abundance, and biomass of key reef organisms as indicators of reef health. CCRE’s assessment program is designed to evaluate similar ecological metrics, so as to be compatible with other monitoring efforts elsewhere in Belize and the western Atlantic Ocean. But researchers have also added some assessments that yield information about key ecological rates and states that are thought to contribute to reef resistance, resilience, and recovery in the face of negative impacts. One example of the measured rates is the grazing rates of herbivores, such as parrotfishes and surgeonfishes, which help keep algae from overgrowing the reef. Researchers also monitor the “states” of the benthic community, such as the health status of important reef-building corals, as well as coral recruitment and growth dynamics. This approach provides more comprehensive ecological monitoring, and informs models of reef dynamics that will be used to generate new insights into reef community structure in response to different reserve management regimes.

This study is designed to take advantage of the strengths and capabilities of the Carrie Bow Cay Field Station and produce important information that will be applied to habitat management in the newly formed SWCMR no-take area.

Research divers survey a permanent transect in the SWCMR.

2018 Scientific Publications Araujo, T. Q. and R. Hochberg. 2017. Description of a new species of Thaumastoderma (Gastrotricha: Macrodasyida: Thaumastodermatidae) from Belize and Tobago. Proceedings of the Biological Society of Washington 130(1), 120–127. https://doi.org/10.2988/17-00003 Ashur, M. M. and D.L. Dixson. 2018. Multiple environmental cues impact habitat choice during nocturnal homing of specialized reef shrimp. Behavioral Ecology, ary171. https://doi.org/10.1093/beheco/ary171 Baker, D. M., C.J. Freeman, J.C.Y. Wong, M.L. Fogel, and N. Knowlton. 2018. Climate change promotes parasitism in a coral symbiosis. The ISME Journal 12(3), 921–930. http://doi.org/10.1038/s41396-018-0046-8 Bornbusch, S.L., J.S. Lefcheck, and J.E. Duffy. 2018. Allometry of individual reproduction and defense in eusocial colonies: A comparative approach to trade-offs in social sponge-dwelling Synalpheus shrimps. PLoS ONE 13(3): e0193305. https://doi.org/10.1371/journal.pone.0193305 Campbell, J. E., A.H. Altieri, L.N. Johnston, C.D. Kuempel, R. Paperno, V.J. Paul, and J.E. Duffy. 2018. Herbivore community determines the magnitude and mechanism of nutrient effects on subtropical and tropical seagrasses. Journal of Ecology 106: 401-412. https://doi.org/10.1111/1365-2745.12862 Cheng, B.S., G.M. Ruiz, A.H. Altieri, and M.E. Torchin. 2018. The biogeography of invasion in tropical and temperate seagrass beds: Testing interactive effects of predation and propagule pressure. Diversity and Distributions 25:285-297. https://doi.org/10.1111/ddi.12850 Dudoit, ‘A., M. Iacchei, R.R. Coleman, M.R. Gaither, W.E. Browne, B.W. Bowen, and R.J. Toonen. 2018. The little shrimp that could: phylogeography of the circumtropical Stenopus hispidus (Crustacea: Decapoda), reveals divergent Atlantic and Pacific lineages. PeerJ 6, e4409. https://doi:10.7717/peerj.4409v Engene, N. , A. Tronholm, and V.J. Paul. (2018), Uncovering cryptic diversity of Lyngbya: the new tropical marine cyanobacterial genus Dapis (Oscillatoriales). Journal of Phycology 54: 435-446. https://doi:10.1111/ jpy.12752 Lesser, M.P., and M. Slattery. 2018. Sponge density increases with depth throughout the Caribbean. Ecosphere 9(12):e02525. https://doi.org/10.1002/ecs2.2525 Marino, C., J. Pawlik, S. López-Legentil, and P. Erwin. 2017. Latitudinal variation in the microbiome of the sponge Ircinia campana correlates with host haplotype but not anti-predatory chemical defense. Marine Ecology Progress Series 565: 53–66. http://doi.org/10.3354/meps12015v

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McMurray S.E., A.D. Stubler, P.M. Erwin, C.M. Finelli, and vJ.R. Pawlik. (2018) A test of the sponge-loop hypothesis for emergent Caribbean reef sponges. Marine Ecology Progress Series 588:1-14. https://doi. org/10.3354/meps12466 Pende, N., J. Wang, P. M. Weber, J. Verheul, E. Kuru, S. Rittmann, N. Leisch, M. S. VanNieuwenhze, Y.V. Brun, and vT. den Blaauwen, S. Bulgheresi. 2018. Host-polarized cell growth in animal symbionts. Current Biology 28(7): 1039-1051.e5. https://doi.org/10.1016/j.cub.2018.02.028 Ross, C., N.D. Fogarty, R. Ritson-Williams, and V.J. Paul, V.J. 2017. Interspecific variation in coral settlement and fertilization success in response to hydrogen peroxide exposure. Biological Bulletin 233(3): 206-218. https://doi.org/10.1086/696215 Studivan, M. S., and J.D. Voss. 2018. Population connectivity among shallow and mesophotic Montastraea cavernosa corals in the Gulf of Mexico identifies potential for refugia. Coral Reefs 37(4), 1183â&#x20AC;&#x201C;1196. http://doi. org/10.1007/s00338-018-1733-7 Toscano, M.A., J.L. GonzĂĄlez, and K. Whelan. 2018. Calibrated density profiles of Caribbean mangrove peat sequences from Computed Tomography for assessment of peat preservation, compaction and impacts on sea level reconstructions. Quaternary Research 89, 201-222. https://doi.org/10.1017/qua.2017.101 Publications with

are open access.

Caribbean Coral Reef Ecosystems | 36

2018 Participants * served as station manager Alanko, Jerry, Tilghman, MD* Alanko, Sandy, Tilghman, MD* Ashur, Molly, University of Delaware, Newark, DE Bardou, Remy, University of California, Los Angeles, CA Beers, Scott, Smithsonian Environmental Research Center, Edgewater, MD* Beers, Lisa, University of California, Berkley, CA* Benson, Maggy, Smithsonian National Museum of Natural History, Washington, D.C. Bock, Megan, Nova Southeastern University, Dania Beach, FL Bohnsack, Karen, Florida Department of Environmental Protection, Fort Lauderdale, FL Borel, Ashley, Lamar University, Beaumont, TX Branson, David, Smithsonian Marine Station, Fort Pierce, FL Brooker, Rohan, Deakin University, Geelong, Australia Brooks, Barrett, Smithsonian National Museum of Natural History, Washington, D.C. Bulgheresi, Sylvia, University of Vienna, Austria Campbell, Justin, Smithsonian Marine Station, Fort Pierce, FL Carlson, Skylar, Smithsonian Marine Station, Fort Pierce, FL Carmack, Olivia, University of Southern California, Los Angeles, CA Carne, Lisa, Fragments of Hope, Placencia, Belize Catching, Katelin, Lamar University, Beaumont, TX Cody, Clyde, Boise, ID Cody, Liz, Boise, ID Cowan, Zara, University of Delaware, Newark, DE Cox, Kieran, University of Victoria, Victoria, British Columbia, Canada Daza, Laura, University of North Carolina at Wilmington, NC Dos Santos, Larissa, Smithsonian Marine Station, Fort Pierce, FL Dramer, Greg, Kalispell, MT* Dramer, Joann, Kalispell, MT* Duckett, Lisa, Smithsonian Environmental Research Center, Edgewater, MD Ellis, Emily, University of California, Santa Barabara, CA Engleman, Abigail, Florida State University, Tallahassee, FL Evans, James, University of North Carolina at Wilmington, NC Farmer, Lucas, Ohio Wesleyan University, Delaware, OH Faux, Victor, Fragments of Hope, Placencia, Belize Feeney, William, University of Queensland, Brisbane, Australia Feller, Candy, Smithsonian Environmental Research Center, Edgewater, MD Finelli, Chris, University of North Carolina at Wilmington, NC 37 | Caribbean Coral Reef Ecosystems

Fogarty, Nicole, Nova Southeastern University, Dania Beach, FL Foltz, Zachary, Smithsonian Marine Station, Fort Pierce, FL Franco, Avelino, Fragments of Hope, Placencia, Belize Frost, Emily, Smithsonian National Museum of Natural History, Washington, D.C. Gaffey, Bethany, Smithsonian Marine Station, Fort Pierce, FL Godfrey, Dale, Fragments of Hope, Placencia, Belize Goodison, Mike, Smithsonian Environmental Research Center, Edgewater, MD Gruhl, Alexander, Max Planck Institute for Marine Microbiology, Bremen, Germany Habicht, Kelly, Nova Southeastern University, Dania Beach, FL Haren, Nick, Anacortes, WA* Haren, Marylin, Anacortes, WA* Harper, Leah, Nova Southeastern University, Dania Beach, FL Hawthorne, Timothy, University of Central Florida, Orlando, FL Henslee, Hayden, Lamar University, Beaumont, TX Hightshoe, Morgan, Nova Southeastern University, Dania Beach, FL Hill, Kate, Florida State University, Tallahassee, FL Hoch, Matthew, Lamar University, Beaumont, TX James, Edwin, Tilgman, MD* James, Bonnie, Tilgman, MD* Johnson, Maggie, Smithsonian Marine Station, Fort Pierce, FL Johnston, Lane, University of Delaware, Newark, DE Jones, Scott, Smithsonian Marine Station, Fort Pierce, FL Joseph, Jennifer, University of Delaware, Newark, DE Kelly, Joseph, Stony Brook University, Stony Brook, NY Kimura, Stephen, University of California, Santa Barbara, CA Koltes, Karen, Garrett Park, MD Lehmann, Michael, St. Louis, MO* Ling, Scott, University of Tasmania, Hobart, TAS, Australia Looby, Audrey, Smithsonian Marine Station, Fort Pierce, FL McMurray, Steven, University of North Carolina at Wilmington, NC McNabb, Mark, University of Belize, Belmopan, Belize Mogg, Andrew, The Scottish Association for Marine Science, Oban, Argyll, Scotland Moore, Joel, Shingle Springs, CA* Moore, Linda, Shingle Springs, CA* Mumby, Peter, University of Queensland, Brisbane, Australia Olinger, Lauren, University of North Carolina at Wilmington, NC Opishinski, Tom, Interactive Oceanographics, East Greenwich, RI Paredes, Gabriela, University of Vienna, Austria Caribbean Coral Reef Ecosystems | 38

2018 Participants Parsons, Keith, Atlanta, GA* Parsons, Shirley, Atlanta, GA* Paul, Valerie, Smithsonian Marine Station, Fort Pierce, FL Pawlik, Joseph, University of North Carolina at Wilmington, NC Penland, Laurie, Smithsonian Scientific Diving Program, Washington, D.C. Peresta, Gary, Smithsonian Environmental Research Center, Edgewater, MD* Pitts, Kelly, Nova Southeastern University, Dania Beach, FL Relyea, Darby, The University of Vermont, Burlington, VT Ritson-Williams, Raphael, California Academy of Sciences, San Francisco, CA Roff, George, University of Queensland, Brisbane, Australia Sauvage, Thomas, Smithsonian Marine Station, Fort Pierce, FL Sayer, Martin, The Scottish Association for Marine Science, Oban, Argyll, Scotland Scheff, George, Boise, ID* Schleiger, Douglas, Smithsonian Scientific Diving Program, Washington, D.C. Segura-Garcia, Iris, Smithsonian Marine Station, Fort Pierce, FL Sherwood, Craig, Deale, MD* Sih, Tiffany, James Cook University, Townsville, Australia Simpson, Lorae, The University of Florida, Gainesville, FLv Smith, Emily, Lamar, Beaumont, TX Steneck, Robert, University of Maine, Walpole, ME Strawhand, Alexander, Florida State University, Tallahassee, FL Trotz, Maya, Fragments of Hope, Placencia, Belize Tschirky, John, American Bird Conservancy, Washington, D.C. Vollmer, Alicia, Nova Southeastern University, Dania Beach, FL Voss, Josh, Harbor Branch Oceanographic Institute, Florida Atlantic University, Fort Pierce, FL Walczak, Joanna, Florida Department of Environmental Protection, Fort Lauderdale, FL Wallen, Mariko, Fragments of Hope, Placencia, Belize Warrender, Tammi, The Scottish Association for Marine Science, Oban, Argyll, Scotland Whippo, Ross, University of Oregon, The Oregon Institute of Marine Biology, Charleston, OR Wulff, Janie, Florida State University, Tallahassee, FL

39 | Caribbean Coral Reef Ecosystems

Acknowledgements Our research is hosted by the Belize Fisheries Department and we thank Ms. Beverly Wade, Mr. Rigoberto Quintana, Mr. Mauro Gongoro, Ms. Felicia Cruz and staff for collaboration and issuing permits. The owners and dedicated staff of Pelican Beach Resort in Dangriga provided logistical support for our fieldwork. Earl David and his staff provided boat transportation, as well as invaluable advice and support.

Numerous volunteer managers helped run the field station and assisted in research activities; we greatly appreciate their many efforts: Jerry & Sandy Alanko, Lisa & Scott Beers, Liz & Clyde Cody, Greg & Joann Dramer, Nicky & Marilyn Haren, Ed & Bonnie James, Mike Lehmann, Joel & Linda Moore, Keith Parsons, Gary Peresta, George Scheff, Lisa Schile, and Craig Sherwood.

In Fort Pierce, we sincerely thank Gloria Baxevanis for administrative advice and assistance with many fund management tasks.

In Washington, Klaus Ruetzler and Mike Carpenter are always willing to share wisdom stemming from their many years of experience in Belize. A number of people at NMNH are always willing to answer questions: Wendy Wiswall, Carol Youmans, Charmone Williams, and Lee Weigt among many others. We also thank the office of the Director of the National Museum of Natural History for continued support.

The CCRE program is supported by Federal funding complemented by the Hunterdon Oceanographic Research Fund.

Photo Credits Cover: Scott Jones page 1: Maggy Benson; page 4: (top to bottom) Zach Foltz, Zach Foltz, Scott Ling; page 5: Scott Ling; page 8: Scott Jones; page 9: Abigail Engleman; page 12: Scott Ling; page 13: Joseph Pawlik; page 15: Janie Wulff; page 16: Scott Ling; Page 17: Nicole Fogarty ; page 20: Timothy Hawthorne; page 21: Raphael Ritson-Williams; page 23: Karen Koltes & John Tschirky; page 24: Karen Koltes & John Tschirky; page 25: Karen Koltes & John Tschirky; page 28: Matthew Hoch; page 29: Syliva Bulgheresi; page 30: Janie Wulff; page 31: Janie Wulff; page 32: Joseph Kelly; page 33: Scott Ling; page 34: Scott Ling; Back cover: Scott Jones

Caribbean Coral Reef Ecosystems | 40

Smithsonian Marine Station Caribbean Coral Reef Ecosystems Program Fort Pierce, FL · Carrie Bow Cay, Belize www.ccre.si.edu www.sms.si.edu